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The Mayfly Family Baetiscidae (Ephemeroptera). Part I Abstract Introduction

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The Mayfly Family Baetiscidae (Ephemeroptera). Part I Abstract Introduction rrc,m: Ar-v/~f·lC·F~-f1 ;;\: ~~;:::; ...':-~--_,!r'·i·:_: :-· 1. :- \ ·:J;c;L()G'f ~·~- ~·., _·· ,, ____ ,_,>-:.. ;-·;.-.;JViarshall {:-·:~-.:nl! ~11 ?ut,;'.shir:g ~::o: ;.:1c.;i· !.:t;0n, 196:0) THE MAYFLY FAMILY BAETISCIDAE (EPHEMEROPTERA). PART I Lewis Berner and Manuel L. Pescador* Department of Zoology, University of Florida Gainesville, Florida 32611 ABSTRACT The history of the taxonomy and phylogeny of Baetisca~ the single genus in the family Baetiscidae, is reviewed beginning with the description of Prosopistoma as a crustacean. The relationship of the Prosopistomatidae to the Baetiscidae is treated as well as the concurrence of the authors with Edmunds' conclusion that the two families are derived from a common ancestor but parallel evolution must explain some of their similarities. There is a summary of presently recognized species and characteristics by which nymphs are differentiated. A brief treatment of the ecology and life history of various species is included. INTRODUCTION If Latreille had had nymphal specimens of Baetisca in 1833, would he have been as puzzled by them as he was by Prosopistoma when he described these curious nymphs as a new genus of crustacean? Certainly, Baetisca nymphs are sufficiently different from other Ephemeroptera to cause one to conclude that he might have done so. The superficial similarities of the mesonotal shields and the gills were enough to lead early entomologists to the misconception that the two are very closely related. Only in recent years, through * Laboratory of Aquatic Entomology, Florida A&M University, Tallahassee, Florida 32307 511 512 LEWIS BERNER AND MANUEL L. PESCADOR modern phylogenetic studies of all stages, has it been recognized that there are significant differences between them. Say (1839) described a new species of Baetisca, which he called obesa, because of its short, robust body. Next, Walker (1853) characterized Baetis obesa both in Latin and English. Hagen (1861) also briefly redescribed the subimago following Say's earlier des­ criptions, and the species was recognized by Walsh (1862) as being a member of his new genus Baetisca after he secured male imagos. In 1863, he remarked (p. 178) "B. obesa Say, Walsh. Both genus and species new to me; very curious. 11 Say's description was based on the subimago. Walsh (1864), after four years, successfully reared adults from the nymph and was then able to relate the two stages. He had sent a specimen of the nymph to Hagen, who wrote to Walsh "The larva No. 66 is the most extraordinary animal I have seen, so that I asked myself whether it really belonged to Insecta. 11 After careful analysis he concluded that it was an insect assigned to the "Ephemerina. 11 Even more surprising, he was able to relate the nymph to an adult Baetisca obesa which Walsh had sent to him earlier. Walsh (1864), with Hagen's confirmation in hand, gave an excellent description of the nymph and discussed his observations of its habitat and emergence. Eaton (1871), in his A Monograph on the Ephemeridae, redescribed the genus and the single, known species, B. obesa in Latin. He illustrated the f orewing (pl. II, Fig. 6) and the male genitalia (pl. V, Fig. 9), both rather poorly. Provancher (1876) reported a species from Quebec as ClBe uni­ color Hagen, but in 1877 decided that his specimens represented a new species, which he named ClBe rubescens~ nov. sp. Subsequently, McDunnough (1921), after studying Provancher's male specimen, assigned it to Baetisca making rubescens the second species to be included in the genus. Joly (1880) published a French translation of Walsh's (1864) description of Baetisca and added comments in footnotes. Here he compared features of Baetisca with those of PI'osopistoma punctifrons (= P. foliaciwn Fourcroy) and P. variegatwn Latrielle. Vayssiere, (1882) in his very comprehensive paper on Ephemeroptera, reported that he saw analogies between PI'osopistoma and Baetisca nymphs in their respiratory apparatus and he discussed their morphology at length as well as providing excellent illustrations. He concluded that Baetisca is to North America what Prosopistoma is to Europe and Madagascar. Eaton (1885) described and illustrated both adult and nymphal stages of Baetisea. Subsequently, Needham (1905) reported that B. obesa had been taken in New York State and in Indiana and he included photographs of the nymph and subi~aginal wings. Ulmer BAETISCIDAE 513 (1920) merely listed the family Baetiscidae with only a single genus, Baetisca, included in it. The next reports of the genus were those of Clemens (1913, 1915) in his study of mayflies of the Georgian Bay area of Lake Huron. Traver (1931) added two new species of Baetisca, carolina and the enigmatic callosa from North Carolina and West Virgina, respectively. Soon afterwards, McDunnough (1932) reported two additional species, B. laurentina and B. Zacustris, from Canada, and also provided a key to the known nymphs. Traver again re­ ported in 1932 on B. carolina in North Carolina and in 1937 she described B. t'homsenae from that State. The latter species was synonyrnized with B. carolina by Berner (1955). The classic work by Needham, Traver, and Hsu (1935), The Biology of Mayflies, gave a comprehensive summary of the taxonomic knowledge of Baetisca. Neave (1934) described B. bajkovi from Manitoba and Spieth (1933, p. 359) concluded that "Baetisca seems a distinct entity in the phylogenetic arrangement of mayflies." He erected the superfamily Baetiscoidea to accommodate the family. Baetisca rogersi was described from northwestern Florida by Berner (1940) and further details were added by him in 1950. He summarized the knowledge of the southeastern species of the genus in 1955 and described two new ones at that time, B. escam­ biensis and B. gibbera. Burks treated the Illinois species in 1953, and in 1962 the late Justin Leonard and his wife Fannie did the same for Michigan. Daggy (1945) reported that the genus occurs in Minnesota. Following World War II, with the great upsurge of interest in aquatic insects, Baetisca was frequently encountered in stream studies and reports of its occurrence grew. In 1959, Berner sununarized our knowledge of the genus to that date in tabular form. Until Edmunds' (1960) report no specie8 was known positively to occur in western North America and in that paper he described Baetisca colwnbiana, a species not subsequently reported. Schneider and Berner (1963) described another Florida species, B. beaki_, from the northwestern part of the State. Later, Pescador and Peters (1971) described the imago of beaki reared from the nymphal stage. The most recently described species, B. bemer1:, is that of Tarter and Kirchner (1978) from West Virginia. The latest treatment of Baetisca was given by Edmunds, Jensen, and Berner (1976) in their book on North and Central American mayflies. PHYLOGENETIC RELATIONSHIPS OF THE BAETISCIDAE Few papers have been published dealing with the relationships of Baetiscidae and Prosopistomatidae. In 1917, Lameere included 514 LEWIS BERNER AND MANUEL L. PESCADOR two subfamilies in his Prosopistomidae, Hexagenitinae, which is composed of the fossil genus Hexagenites, and Prosopistominae, divided into two tribes, the Baetiscini and Prosopistomini. In a discussion of the systematic position of Prosopistoma, Gillies (1954) concluded that both Baetisca and Prosopistoma are derived from a common stock of great antiquity, but since then evolution and specialization have proceeded independently. His conclusion was based on the nymphs having many structural features in common, notably a mesonotal carapace, branchial chamber, and cephalization of the central nervous system, and the adults having the similarly unique arrangement of the cubital veins of the wings. Fontaine (1958), studying the affinities of Prosopistoma, noted the nymphal similarities with Baetisca. She proposed two causes that might result in the similarities: (1) FTosopistoma and Baetisca possess true systematic affinities and come from the same ancestor and (2) FTosopistoma and Baetisca are two evolved genera--PY'osopistoma having the higher degree of evolution (the mesonotal shield forming a true respiratory chamber, the labium completely encircled in its basilar portion, the nervous system with the thoracic and abdominal ganglia fused into a single mass). They are parallel evolutionary lines and are distinct from the re­ mainder of the Ephemeroptera. Fifty-two years after the appearance of his study of the morphology of immature Ephemeroptera, Vayssiere published in 1934 an anatomical study of Baetisca obesa and B. carolina. He con­ cluded that the internal anatomy of Baetisca is much like that of PY'osopistoma and that his ideas put forward in 1882 about the re­ lations of the two had now been authenticated. Landa's (1969) investigation of the comparative anatomy of mayfly nymphs indicated that the Neoephemeridae showed close re­ lationships to the Caenidae, Prosopistomatidae, and particularly to the Baetiscidae. In the Caenidae, he concluded that Brachycercus is more closely related to Baetisca than is Caenis. His phylogeny, based on the comparative anatomy of the tracheal system, Malpighian tubes, and the nerve band of nymphs and illustrated in a 1973 paper, represented the Neoephemeridae and the Baestiscidae as being related and the Caenidae related to the Prosopistomatidae. The four families are derived from a common stem. Demoulin (1956 and 1969) presents cogent arguments in support of his contention that the Baetiscidae are derived from the Onis­ cigasterinae and the Prosopistomatidae from the Ameletopsinae and that both families should be included in t.he superfamily Siphlonu­ roidea. He has given an interesting analysis of morphological evidence leading him to his conclusions; however, we remain in agreement with Edmunds' interpretation of the phylogeny of these BAETISCIDAE 515 [J< Baetisca obesa (Say) ~ Baetisca lacustris McD. m Baetisca laurentina McD. ~-- .. ~. Figure 1. Presently known distribution of the species of Baetisca in North America. two families. In Edmunds' comprehensive study (1972) of the evolution of Ephemeroptera, a diagram of a probable phylogeny for the families and subfamilies was given.
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